北海道大学苫小牧研究林に見られる樹上性および土壌性ダニ類(ダニ目:トゲダニ亜目、ササラダニ亜目)
誌名誌名 北海道教育大学紀要. 自然科学編
ISSNISSN 13442570
巻/号巻/号 581
掲載ページ掲載ページ p. 23-36
発行年月発行年月 2007年8月
農林水産省 農林水産技術会議事務局筑波産学連携支援センターTsukuba Business-Academia Cooperation Support Center, Agriculture, Forestry and Fisheries Research CouncilSecretariat
北海道教育大学紀要(自然科学編)第58巻 第 1号
Journal of Hokkaido University of Education (Natural Sciences) Vol. 58, NoJ
Arboreal and Forest Floor Mites (Acari: Gamasida, Oribatida)
Found in the Tomakomai Experimental
Forest of Hokkaido University, Hokkaido, Northern ]apan
TAKAKU Gen and SASAKI Tomiya
Department of Biology, Sapporo Campus, Hokkaido University of Education, Sapporo 002-8502
北海道大学苫小牧研究林に見られる樹上性および土壌性ダニ類
(ダニ目:トゲダニ亜日,ササラダニ亜日)
高久 元・佐々木富也
北海道教育大学札幌校生物学教室
ABSTRACT
平 成 19年 8月August. 2007
This study lcioks at oribatid and gamasid fauna of arboreal habitats on the oak Quercus mongolica
var. grosseserrata that has been enclosed by a scaffolding system as well as mite fauna of the forest floor
surrounding an oak tree adjacent to the scaffolding system in cool-temperate forest in the Tomakomai
Experimental Forest of Hokkaido University, Hokkaido, northern Japan. As part of this study we col-
lected a total of 2,596 individuals from 61 species of gamasid mites, including 6 species recorded in J apan
for the first time, aI}d 8,640 individuals in 84 species of oribatid mites, Most of the collected gamasids and
oribatids were ground dwellers and inhabited the forest floor, although we did collect several species
that were only found in the bark or leaves in the forest canopy. Sorensen's coefficient index and com-
plementarity between each site indicated that the mite fauna located on bark and leaves were actually
distinct from forest floor fauna. These results suggest that species specific to bark and leaves utilize
these as a permanent habitat. We also found that two species of the family Phytoseiidae could only be
found in the canopy leaves and 3 species of oribatid mites primarily found on bark surfaces were domi-
nant at a particular level of the trunk and canopy. We believe this specificity to certain heights and areas
may be caused by a distribution of resources (prey and organic matter), bark structure, or abiotic fac-
tors on bark surface, although additional study is necessary for elucidating these issues further.
INTRODUCTION
Mites (Acari) are one of the most diverse and abundant groups among the terrestrial arthropods,
23
高久 元 ・佐々木富也
with over 40,000 named species of mites and estimates for the total number of species rise to anywhere
between 500,000 to one million species (Walter and Proctor, 1999). The principal habitat of mites consists
of soil and litter, and these areas generally have the greatest diversity and abundances of mites (Walter
and Behan-Pelletier, 1999). 1n addition to soil and litter, species-rich mite assemblages are also found on
bark and in forest canopies. Most of the biodiversity studies that focus on the forest canopy, however, have
so far tended to focus on insects, and mites have largely been ignored or grossly underestimated (Walter,
1993; Walter and Behan-Pelletier, 1999). Recent years, however, have seen this oft-overlooked animal from
forest canopies and tree trunks being investigated in subtropical. tropical and temperate forests, with these
studies revealing that several different kinds of mites inhabit the arboreal environments (Walter, 1993;
Watanabe, 1997). Several papers have already reported that arboreal fauna is distinct from that of soil and
litter (Aoki, 1973; Ito, 1986; Walter et al., 1994; Winchester, 1997; Proctor et al., 2002), whereas Beaulieu et al.
(2006) suggested that tree trunks represent a highway for dispersal between habitats for most mesos-
tigmatic mites that use them, and Watanabe (1997) noted that mites found on tree trunks or in the canopy
usually migrate from the soil substratum. Aoki (1973) divided oribatid mites into 3,groups -soil forms,
wandering forms, and arboreal forms -and speculated that some of these mites are restricted in the soil or
arboreal environment, while some wander between the tree canopy and the soil.
Most of the above-mentioned results are based on surveys carried out in tropical. subtropical, and
temperate forest, and to date there has been no comprehensive study on the arboreal mites in cool-
temperate forest that is found in Hokkaido, northern J apan, except for a study on arboreal arthropod
fauna in the Daisetsu mountain area of central Hokkaido (Yamashita and 1shii. 1976). The majority of stu-
dies on canopy mites in Japan have tended to focus on oribatid mites and have been carried out on con-
ifers (Karasawa and Hijii, 2005), meaning that information regarding gamasid mites in forest canopies on
broad-leaved trees remains relatively scarce, although one exception to this have been studies of phy-
toseiid mites as the natural enemy of
Fig. 1. A. Scaffolding system in Tomakomai Experimental Forest of Hokkaido University; B. Tullgren funnel for col-
lecting mites from litter and soil samples; C. Bark spraying method for collecting mites from bark surface.
24
北海道大学苫小牧研究林に見られる樹上位および土壊性ダニ類 (ダニ目:トゲダニ亜日,ササラダニ亜日)
pare mite species that simu1taneous1y inhabit the ground. tree trunk. and canopy (Ito. 1986; Nadkarni
and Longino. 1990; Fagan et al.. 2006). In the present study. we 100ked to investigate oribatid and gama-
sid fauna of arborea1 habitats on the oak tree (deciduous broad-1eaved tree) and forest floor surrounding
the trees in the coo1-temperate forest in Tomakomai Experimenta1 Forest be10nging to Hokkaido Uni-
versity. Hokkaido. notthern Japan. This study a1so compares fauna between samp1ing sites. We a1so dis-
cuss issues such as species richness. distinctiveness. site specificity and the distributiona1 pattern of
arhorea1 and ground-dwelling oribatid and gamasid mites.
MATERIALS AND METHODS
Research area and foca1 tree species
All samp1ing was carried out in coo1-temperate forest in Tomakomai Experimenta1 Forest of Hokkaido
University (420
40'N. 1410
141'E). Hokkaido. northern Japan. The experimental forest is 10cated in Tomako-
mai City. approximate1y 60km south of Sapporo City. All mite samp1es were collected from 1eaves and bark
of the oak Quercus mongolica var. grosseserrata that had been enclosed by a scaffo1ding system (about 15m
x 15m in width. 21m in height; Fukushima et al.. 1998) (Fig. 1A) set in the experimenta1 forest and from
1itter and soi1 of the forest floor surrounding di宜erentoak tree adjacent to the sca宜Oldingsystem.
Collection
All samp1es used in the present study were collected on 20 Ju1y. 29 August. 4 October. 1 November
2005. 15 May and 26 June 2006. Samp1ing of mites from the soil. 1eaf 1itter. bark surface. and 1eaves was
carried out using the following methods.
1) Samp1ing from soil and litter
A 10m x 10m p10t was estab1ished on forest floor surrounding an oak tree adjacent to the sca旺old-
ing system. The p10t was then further subdivided into 25 subp1ots. each 2m x 2m in size. In order to co1-
1ect 1eaf litter samp1es. 10 subp10ts were chosen random1y from 25 subp10ts on each research date. and
1itter from a 20cm X 20cm area was taken at each chosen subp1ot. Every effort was made to avoid in-
cluding any of the top 1ayers of the soi1 as part of the 1itter samp1es. Mites were extracted from the litter
samp1es over 3 days using Tullgren funne1s with 20W light bu1bs (Fig. 1B). Soi1 samp1es were collected
from random1y se1ected l() subp10ts by using a cubic samp1er (5cm x 5cm x 5cm) on each research
date. As far as possib1e. 1eaf 1itter was removed from the samp1ing subp10t before collecting soil samp1es.
The method used on extracting mites from soi1 samp1es was the same as for the litter samp1es.
2) Samp1ing from bark surface
In an attempt to collect mites from the bark surface of oak trees. bark spraying method was em-
p10yed (Nakashizuka and Stork. 2002). The collecting hoop. with a p1astic screw-top container containing
70% ethano1 at the base of the hoop. was pinned tightly to the bark using thumbtacks. A O. 5m x O. 5m
area of bark above the edge of the hoop was sprayed thorough1y using commercia1 pyrethroid insecti-
cide. After approximate1y 20 minutes. the sprayed area was brushed gently using a paintbrush and a
number of samp1es ・thatincluded mites -were gathered in a container (Fig. 1C). In carrying out the
samp1ing. every effort was made to avoid parts of the bark covered with 1ichen and moss. Samp1ing of
25
高久 元・佐々木富也
bark surface was carried out at heights of 5m, 10m, and 15m from the groundusing the sca宜'oldingsystem.
3) Sampling from leaves
The sampling of leaves from the oak canopy was carried out at heights of 5m, 10m, and 15m from the
ground using the sca旺'oldingsystem. At each sampling height, 50 leaves were randomly chosen, cut from
branches, and then placed into sealable plastic bags. Mites were directly separated from leaves using for-
ceps and a fine paintbrush under the stereoscopic microscope, and then五xedin 70% ethanol.
Regarding the method for collecting arthropods from the forest canopy, canopy knockdown, or insecti-
cide fogging, is a well-known and popular method (Nakashizuka and Stork, 2002; Yanoviak et al., 2003) .
This technique is e妊icientin surveying arthropods living on the inaccessible forest canopy. The canopy
does, however, include several di旺'erentmicrohabitats, such as leaves, bark crevices, suspended soil. lichen,
and moss. Even if it were possible to collect canopy mites by canopy knockdown, it would still be difficult
to precisely identify their habitat. Moreover, Yanoviak et al. (2003) have shown that many arthropods that
are killed by insecticide remain trapped within the epiphyte material, and Watanabe (1997) indicated that
repeated sampling of the same sites by insecticide smoking is di伍cult.As the present study is expected to
clarify mite fauna in each research site and specificity to certain habitats through repeated sampling, we
therefore decided to employ the direct method in collecting mites from bark surface and leaves.
All mite specimens were cleared in 60% lactic acid or Nesbitt's solution (Krantz, 1978). Gamasid
mites were mounted in Hoyer's medium on glass slides and oribatid mites were mounted in lactic acid on
glass slides. The mite specimens were identified to species as far as possible, and this was performed by
differential interference contrast microscope.
Similarity and complementarity of fauna between research sites
In this study we used Sorensen's coe妊icientindex for indicating similarity of mite fauna between re-
search sites. Sorensen's coe伍cientindex is regarded as one of the inost e旺ectivepresence/absence similar-
ity measures (Magurran, 2004). The index (Cs) was ca1culated as follows: Cs = 2a / (2a + b + c) : a, the
total number of species present in both sites; b, the number of species present only in one site; c, the num-
ber of species present only in the other site. The index ranges from zero (no overlap) to 1 (same species
composition) .
We also ca1culated the Marczewski-Steinhaus distance (Magurran, 2004) as a measure of com-
plementarity in ground vs. canopy mite fauna in comparing these with results of any previous studies. We
calculated complementarity (CMs) is ca1culated as follows: CMs = 1-a / (a + b + c) : variables that are
the same as for Sorensen's coefficient index. The complementarity ranges from zero (same species com-
position) to 1 (no overlap).
Site specificity
In order to clarify the site specificity of mites, each mite species was assigned to one of the following
4 categories: 1) Ground dweller: Species collected from only forest floor (soil and litter) ; 2) Bark spe-
cialist: Species exclusively collected from bark; 3) Foliar specialist: Species primarily found on leaves;
4) Generalist: Species collected from both forest floor and bark.
To verify whether species collected from bark and leaves were distributed among a certain height of
26
北海道大学苫小牧研究林に見られる樹上性および土壌性ダニ類(ダニ目:トゲダニ亜日,ササラダニ亜日)
Table 1. Abundance of the gamasid mite species collected from each study site (all sampling dates are pooled) .
七speciesrecorded from ]apan for the first time.
27
Q凶
-E
一00
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0
0
0一00
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Species name
*Neogamasus davydovae Volonikhina. 1993 *Neogamasus insularius Volonikhina. 1993 Neogamasus sp. 1 Neogamasus sp. 2 Neogamasus sp. 3 Parasitidae sp Gamasolaelaps ctenisetiger Ishikawa. 1978 Ve羽田iakawasawai Ishikawa. 1982
*Veigaia ochracea Bregetova. 1961 Veigaia uenoi Ishikawa. 1972
竺盟主旦*DendrolaelaJ問。rmatu坦 Hirschmann.1960
D'endrolaelapsル叫oaeIshikawa. 1977 Gama.ゆhissp目
Antennoseius j砂開icusIshikawa. 1979 Asca aPhidioides (Linnaeus. 1758)
Asca nubes Ishikawa. 1969 Asca sculptrata Aoki. 1968 Lasioseius sugawarai Ehara. 1964 Lasioseius初mokoaeIshikawa. 1969 Lωioseius sp. Leioseius brevisetosus Ishikawa. 1969 Leio四eiussp. near insignis Hirschmann. 1963 Z'erconopsis sinuata Ishikawa. 1969
'*Neozercon insularis Petrova. 1977 Zercon armatus Aoki. 1966
Zercon jajうonicusAoki.1964 Zerconidae sp Amblyseius ishizuchiensis Ehara. 1972 Amblyseius orientalis Ehara. 1959 Euseius finlandicus (Oudem加 s.1915)
Euseius sojaensis (Ehara. 1964)
Neoseiulus rePletus (Wu & Li. 1985)
Neoseiulus sp. near haimatus (Ehara. 1967) Phytoseius {Dubininellus) blakistoni Ehara. 1966 T'yphlodromus (Anthoseius) vul.加risEhara. 1959 E:picriopsis stellata Ishikawa. 1972 Podocinum aokii Ishikawa. 1970 P'odocinum catenum Ishikawa. 1970 E:picrius nemorosus Ishikawa. 1969 Macrocheles se門官tusIshikawa. 1968 Gamasholaspis akimotoi (Ishikawa. 1966)
Gamasholaspis browningi (Bregetova & Koroleva. 1960)
Parhola.ゆulusalstoni Evans. 1956 Parholasρulus arboreus Ishikawa. 1980 hァho加;pulω communisOshikawa. 1966)
均時olaspulωde仰 tusIshikawa. 1969 Parhola.ψulus ochraceus (Ishikawa. 1966)
品時ol,時 ulussp. *時ρoω;Pisvacua (Michael. 1891)
正{ypoasPisqueenslandicus (Womersley. 1956) Hypoas.ρis sp. near claviger (Berlese. 1883) 互盟盟企旦:lVeoparasitusρunctatωIshikawa. 1987 Dinychus kurosai Hiramatsu. 1978 Uropodidae sp. 1 Uropodidae sp. 2 Uropodidae sp. 3 Uropodidae sp. 4 Uropodidae sp. 5 Uropodidae sp. 6 Larvae Nymphs Undetermined
Family
Parasitidae
Superfamily
Parasi toidea
Veigaiidae
Digamasellidae
旦!盟主旦asidaeAscidae
Zerconidae
Rhodacaroidea
Ascoidea
Phytoseiidae Phytoseioidea
Ameroseiidae Podocinidae
白区坦竺Macrochelidae Parholaspidae
Eviphidoidea
Laelapidae Dermanyssoidea
Unknown Uropodidae Uropodoidea
Total
元・佐々木富也高久
Cymbaeremaeoidea Cymbaeremaeidae
28
QAM
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山
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Species name
Brachychthonioidea Brachychthoniidae
Family
Hypochthoniidae
Eniochthoniidae
Mesoplophoridae
Lohmanniidae
Eulohmanniidae
Epilohmanniidae
Table 2. Abundance of the oribatid mite species collected from each study site (all sampling dates are pooled) .
Abundance
Litter Bark
17 0
4 0 586 0
185 0
206 0
7 0
o 0 o 0
o 0
o 0
o 2 3
o 0 36 0 o 0 38 0
67 2 o 0
20
17 0
66 2
26 0 5 0
2 0
5 0
o 0
33
8 0 o 0
o o 0 o 11
9 0
o o 9
o o
36 0
o 744 0
34 0
34 0
o 162
7 0
o o
2 0
6 0
39 7
o o 0 2 0
10 0
210 0
4
o 0
3 0
26 0
o 0 13 0
o 2 3
o 19
必一1
1一ωη一お一日一日一お一
7
5一0
0
3
m
1一位一μ一9
0・1一目印一お
7
U一ロ一O一O刀
2
1
1
U
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W
O一1一O一O
O一3日一
I
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一
Liochthonius strenzkei Forsslund. 1963 Poecilochthonius spiciger (Berlese, 1910)
Eohypochtho叩iusmagnus Aoki. 1977
的ρochthoniusrufulus C. L. Koch, 1836
Hアlうochthoniellaminutissima (Berlese, 1904)
M'esoPloPhora (Partlophora) jφonica Aoki. 1970
hrpillacarus hirsutus (Aoki. 1961)
Eulohmannia ribagai Berlese. 1910
Epilohmannia spathulata Aoki. 1970
E:pilohma明nωsp.Phthiracarus bη'obius Jacot. 1930
Phthiracarus japonicus Aoki. 1958
Phthiracarus setosus (Banks, 1895)
Phthiracarus sp. 2
P'hthiracarus sp. 3 Atropacarusはtropscarus)striculus (c. L. Koch, 1836)
Rhysotritia ardu沼(c.L. Koch. 1841)
Camisia la.仲間ica(Tragardh. 1910)
Ca1到isiasegnis (Hermann. 1804)
P'latynothrus yamasakii (Aoki, 1958)
Malaconothrus jψonicωAoki.1966
Alalaco叩othrusPygl珂aeusAoki. 1969
Nothrus biciliatus C. L. Koch. 1841 Nothrωρalustris C. L. Koch. 1839
N'othru沼 silvestrisNi∞let. 1855
IVanhermannia elegantula Berlese. 1913
Hexachaetoniella japonica (Aoki & Suzuki. 1970)
Belba sp. 1 Belba sp. 2
Belba sp. 3
Epidamaeus a伐'gulatusFujikawa & Fujita. 1985
Epidamaeus grandjea百iBulanova-Zachvatkina. 1957
Damaeidae sp. 1
Damaeidae sp. 2
C'epheus ceρheiformis (Nicolet. 1855)
CaenosamerusゆatiosusAoki. 1977
Fosse月間usquadripertitus Grandjean. 1965
Eremobelba jゆonicaAoki. 1959
Eremulus avenifer Berlese. 1913
C'ultroribula lata Aoki, 1961
Ceraω妙 iabipilis (Hermann. 1804)
CeratopPia quadridentata (Haller. 1882)
C'eratoppia sp. X'enillus' tegeocranus (Hermann. 1804)
c'arabodes peniculatus Aoki. 1970 Fissicepheus clavatus (Aoki. 1959)
FissicePheus sp TectoceJうheuscuゅidentatusKnulle. 1954
T'ectoceρheus velatu沼 (Michael.1880)
Lauro虫piadecempectinata (Fujikawa. 1986)
Medioxyoppia sp.
Microρρia minus (Paoli. 1908)
Multioppia (Multilanceゆかa)brevipectinata Suzuki. 1975
0ρ,'piella nova (Oudemans. 1902)
Quadroppia qωd門 carinata(Michael. 1885)
Ramusella (Ramusella) sengbuschi Hammer. 1968
S'ubiasella sp
Flagrosuctobelbella naginata (Aoki. 1961)
Suctobelbella tuberculata Aoki. 1970
SUctobelbella sp. near longisensillata Fujita & Fujikawa. 1987
Ametroproctus (Coropoculia)陀 ticulataAoki & Fujikawa. 1972
ScaPheremaeus yamashitai Aoki. 1970
Licneremaeus novaeguineae Balogh. 1968
Hypochthonioidea
Eniochthonioidea
Mesoplophoro,idea
Lohmannioidea
Eulohmannioidea
Epilohmannioidea
Superfamily
Phthiracaridae
Steganacaridae
Euphthiracaridae
Camisiidae
Nanhermanniidae
Pedrocortesellidae
Damaeidae
Xenillidae
Carabodidae
Otocepheidae
Malaconothridae
Cepheidae
Ameridae
Damaeolidae
Eremobelbidae
Eremulidae
Astegistidae
Metrioppiidae
Nothridae
Ph thiracaroidea
Euphthiracaroidea
Crotonioidea
Nanhermannioidea
Gymnodamaeoidea
Damaeoidea
Liacaroidea
Cepheoidea
Eremuloidea
Carabodoidea
Tectocepheidae
Oppiidae
Tectocepheoidea
Oppioidea
Suctobelbidae
Licneremaeidae Passalozetoidea
北海道大学苫小牧研究林に見られる樹上性および土壊性ダニ類(ダニ目:トゲダニ亜日,ササラダニ亜日)
QO
一
叩
200一00
0
0
0一OO一O一O一O一OO一OO一O一OO一O一O一00
0一0
9u e
yb
Abundance L,itter Bark
11 0
o 2 o 24 o 11
1 0 o 2
o o
o 1
望一一」1 0
o 0
5 0
o 0 2 0
o 0 148
O.
o 4
2 0
3 0
21 0
1165 223 3 1
3847 541
叫一
1
0一00
0
0
0一OO一O一O一5一m
l一01一回一
0
0一O一3-mU2一位
s
-
1
1
一2
-A
一auτ
Table 2. (comtinued)
Incabates angustus Hammer. 1967 I'ncabates .major Aoki. 1970 Eρoribatula sp. near tuberosa Fujikawa. 1972 Oribatula sakamorii AokL 1970 Oribatula sp あIgoribatulatrucata Aoki. 1961 z.ygoηbatula sp. Oripoda sp. near ρinicola Aoki & Ohkubo. 1974 T'runcopes sp. near optatusωiaticus Aoki & Ohkubo. 1974 1Veoribates roubali (Berlese. 1910) Transo門 batessp. near agricola (Nakamura & Aoki. 1989) Scheloribates latipes (C. 1. Koch. 1841) Xylobates loPhothrichu沼 (Berlese.1904) Xylobates sp αratozetella imperatoria (Aoki, 1963) C'eratozetidae sp. c'hamobatesρωillω(Berlese. 1895) Eupeloρs acromios (Hermann. 1804) Eゆeloρ'ssp. near acromios (Hermann.1804) Oribatella sp. T'richo,伊 lumnanゆ onica(Aoki. 1966) Larvae Nymphs Undetermined
Species name
Haplozetidae
Oribatulidae
Family Superfamily
Oripodoidea
Oripodidae
Parakalummidae Prot刀ribatidaeScheloribatidae Xylobatidae
Ceratozetidae Ceratozetoidea
Chamobatidae Phenopelopidae
Oribatellidae Ga1umnidae
Phenopelopoidea
Oribatelloidea Galumnoidea
Total
Table 3. Number of mite species and abundance of adult mites Cin parentheses) collected from each study site.
Total Leaves Bark Litter Soil
61(1. 794) 5 (381) 3 (19) 41 (848) 43(546) Gamasida
84(6.068) 0(0) 19 (317) 58(2.658) 54(3,093) Oribatida
tree, we carried out a chi-square test in a few species of gamasids and oribatids that primarily inhabited
among bark and leaves.
RESULTS
We were able to collect a total of 2,596 individuals Gncluding 802 immature and undetermined spe・
cimens) in 61 species of gamasid mites belonging to 14 families from sampling at 4 sites (Table 1). This
includes 6 species belonging to 5 families that were recorded in ]apan for the first time. Regarding the
oribatid mites. we collected 8,640 individuals Gncluding 2,572 immature and undetermined specimens)
in 84 species belonging to 43 families (Table 2). Most of the collected gamasid and oribatid species were
recorded from soil or litter, whereas species richness and abundance in bark and leaves were relatively
low (Table 3). Although we did not collect any oribatid mite from leaves, species richness and abundance
of oribatid mites were both much higher than those of gamasid mites in the other 3 sites. The character-
istics of fauna and dominant species or family in each site were as follows.
1) Soil: Among gamasid mites, the families Parholaspidae and Uropodidae were dominant, with over
29
高久 元・佐々木富也
68% of adult specimens collected from soil. In oribatid mites, approximately 38% of adults collected from
soil were made up of the species of the family Oppiidae, with Opρiella nova being the dominant species.
More than 100 individuals were recorded within several different oribatid species.
2) Litter: We found that over half (53%) of adult gamasid mites collected from leaf litter were
members of the family Ascidae. Among oribatids, Cultroribula lata (Astegistidae) and Eoh~ρochthonius
magnus (Hypochthoniidae) formed 50% of adult specimens.
3) Bark: We collected 4 gamasid species -Asca nubes, Lasioseius sp. (Ascidae) ,Amblyseius orienta-
lis (Phytoseiidae), and Parhola~ρulus arboreus (Parholaspidae)・frombark, but they had a relatively low
abundance of under 10. In contrast to these findings, 19 species of oribatid mites were recorded from
bark, with Ceratoρρia sp. (Metrioppiidae) comprising over half (51%) of the adult oribatids collected
from bark.
4) Leaves: Here, Euseius finlandicus (Phytoseiidae) was dominant and occupied 83% of gamasid
mites collected from leaves. All species collected from leaves belonged to the family Phytoseiidae. We
were unable to find any oribatid mites on leaves during the present study.
Table 4. Sorensen's coefficient index (upper) and number of species in common (below) between study sites,
based on species composition of gamasid mites.
Leaves Bark Litter
Leaves 0.22 0.04
Bark 1 0.04
Litter 1 1
Soil O 1 29
Table 5. Sorensen's coefficient index (upper) and number of species in common (below) be-
tween study sites, based on species composition of oribatid mites.
Bark
Litter
Soil
Bark
9
6
Litter
0.23
37
Soil
0.16
0.66
Soil
O
0.04
0.69
Table 6. Oribatid species complementarity in canopy vs. ground (data cited from Behan-Pelletier and Walter. -
2000; Proctor et al., 2002), and the complementarity of gamasid and oribatid species in the present study.
Location Canopy microhabitat
Germany Canopy twigs and leaves; bark
Canada Canopy mosses
Central ]apan Canopy branches, mosses, lichens
Australia Bark
Northern ]apan: Oribatida (present study) Bark
Northern ]apan: Gamasida (present study) Bark,leaves
30
Complementarity
0.42
0.68
0.85
0.97
0.88
0.98
北海道大学苫小牧研究林に見られる樹上性および土壊性ダニ類 (ダニ目:トゲダニ亜日,ササラダニ亜日)
Similarity of fauna between research sites
Sorensen's coefficient index and the number of common species found between study sites were
highest between soil and litter fauna in both Gamasida and Oribatida (Tables 4. 5). The index and the
number of common species between ground and bark or leaves in Oribatida and Gamasida were both
much lower than for those values between soil and litter. Complementarity of species composition of
gamasids and oribatids between ground and canopy in the present study were 0.98 and 0.88. respective-
ly. These figures are similar to or greater than any previously reported results (Table 6). indicating that
mite fauna found on bark and leaves are distinct from forest floor fauna.
Site specificity
In this study we found most gamasid and oribatid mites to be ground dwellers and associated with
soil and leaf litter on the forest floor (Table 7). A small portion of gamasids were specific to canopy bark
or leaves. and generalists. commonly occurring arboreal habitat and forest floor. were rare. A small range
of oribatids was represented by bark specialists (11%) and generalists (12%). Most generalists were pre-
dominantly found in one site. such as either in the forest floor or canopy; for example. Hexachaetoniella
JfJ.ρonica (Pedrocortesellidae) was collected from bark with the exception of 1 specimen taken from leaf
litter. and Op.ρiella quadricarinata (Oppiidae) was collected from the forest floor with the exception of 1
specimen that was found in bark.
Chi-square test was performed on 2 species of the family Phytoseiidae that were only found on
leaves in the canopy and 3 species of oribatid mites that dominated the bark surface area. The results of
the test indicated that distribution of every species di百'eredsignificantly based on height. Euseius finland-
icus tended to be found on leaves in lower areas that were 5m and 10m in height. while Typhlodromus
(Anthoseius) vulgaris was found in higher areas (15m) of the canopy (Table 8). The oribatid mite spe-
cies Ceratop'ρia sp. was mainly distributed on leaves in lower areas. whereas Hexachaetoniella jゆonica
andEρoribαtula sp. near tuberosa were found in the higher parts of the canopy (Table 9).
Table 7. Number of sp~cies of ground-dweller. bark specialist. foliar specialist. and generalist in gamasid and oriba-
tid mites (percentage of each category in parentheses indicates relative abundance) .
Ground-dweller Bark specialist Foliar specialist
Oribatida
53 (87%)
65(77%)
2(3%)
9(11%)
5(8%)
0(0%)
1(2%)
10(12%)
凶
一1
4
h
一68
Generalist
Gamasida
Table 8. Number of phytoseiid mites Euseius finlandicus and TYPhlodromus (Anthoseius) vulgaris collected from
leaves at each height.
5m 10m 15m
Euseius finlandicus 144 118 56 *** TYPhlodromus (Anthoseius) vulgaris 4 5 28 *** *本*
: p < o. 001(chi-square test)
31
高久 元・佐々木富也
Table 9. Number of oribatid mites Ceratoρρia sp・,Hexachaetoniella jゆonica,and Eporibatula sp. near tuberosa col-
lected from bark surface at each height.
5m 10π1 15m
Ceratoρρia sp. 65 68 29 *** Hexachaetoniella jゆomca 1 2 30 *** Eρoribalula sp. near tuberosa 1 9 14 ** 料*:p <0.001;紳:p <O.Ol(chi-square test)
DISCUSSION
This study revealed that species richness of gamasid and oribatid mites on the ground (soil and lit-
ter) was far higher than that of the canopy (bark and leaves). This lower diversity in the canopy may
be caused by a corresponding lower availability of resources. and total organic matter is often the most
important factor influencing the abundance and community structure of soil Collembola and Oribatida
(Hasegawa. 2001). Organic matter may also affect fungal biomass, which is consumed by some mites.
Although little data shows di旺erencesbetween canopy and forest floor organic matter. they di宜ergreat-
ly in a variety of physical and chemical characteristics (Nadkarni and Longino. 1990). and both chemistry
of organic matter and decomposition rates have an important e百ecton the trophic relationship and com-
munity structure in soil arthropods (Berg et al.. 1998). Abundance and/or quality of organic matter in
the canopy may determine the abundance and diversity of mites in the canopy. Greater abiotic extremes
in the canopy may also result in lower diversity and abundance of mites in the canopy. Tree crowns are
subjected to more extreme variability in ambient insolation, wind. desiccation, and precipitation than
found in the forest understorey (Bohlman et al. 1995; Fagan et al.. 2006). Such extreme variability and
other environmental conditions a百ectthe diversity and abundance of mites in the canopy.
The majority of gamasid and oribatid species collected in this study were ground dwellers, inhabit-
ing the forest floor. Some were specific to bark or leaves. however. and complementarity between ground
and canopy mite fauna in both gamasids and oribatids was greater here than in the results of previous
studies. A study by Watanabe (1997) on arboreal arthropods collected by insecticide smoking concluded
that there is no distinct di百erencebetween arboreal and ground-dwelling oribatid mites. which depend
mainly on dead plant debris. in temperate forests. A number of studies. however. have revealed that
most arboreal mites, especially oribatids. distribute exclusively in their habitats (Ito, 1986; Behan-Pelletier
et al., 1993; Walter et al.. 1994; Walter and Behan-Pelletier. 1999). Some gamasid mites may also use tree
trunks as a permanent habitat, although most gamasids use the trunk as a highway for dispersing be-
tween habitat patches (Beaulieu et al.. 2006). Bark and foliar special
32
北海道大学苫小牧研究林に見られる樹上性および土様性ダニ類.(ダニ目:トゲダニ亜目,ササラダニ亜日)
floor and canopy habitat. Most generalists were, however, found predominantly in one site and rarely
found in the other habitat. Whether these generalist species simply utilize the bark for dispersing or
actually use these areas as a habitat for feeding and reproducing remains unknown. The canopy mites
found on the ground may be as a resurt of random suspended soil or litter fall, while the forest floor
mites found in canopy habitat may occasionally use the canopy as a temporary habitat.
In the present study, we were unable to collect any oribatid mites from canopy leaves. Although
data obtained from cool-temperate forest is insufficient for discussing why there were no oribatids on
leaves, there may be some plausible explanations for this result:
(1) A difference in sampling methods: Karasawa and Hijii (2005) indicated that the washing method,
in which the mites were washed off leaves using a solution of water mixed with detergent, was more
likely to be an effective and a relatively unbiased method for collecting oribatid mites from the canopies
of broad-leaved trees in ]apan. The difference in sampling method may yield a different result between
the present study and previous ones. In their paper, however, the authors also tried to collect oribatids
using the direct method, in which the mites were directly collected from leaves using a brush or forceps,
and recorded nearly 300 individuals of 5 oribatid species from the leaves of broad-leaved trees. In addi-
tion to this particular study, Walter et al. (1994) used the direct method to record 28 oribatid mite spe-
cies from the leaves, stem, and trunk of the musk daisy in temperate forest located in Victoria, Australia,
as we performed in the present study. The resu1ts of. the above-mentioned studies indicate that the
direct method is not entirely useless in collecting mites from the leaves of broad-leaved trees, and the
sampling method should not affect the result of the present study.
(2) Di旺erencesin climatic province: Studies on canopy mites performed in tropical. subtropical. and
temperate regions, have recorded many kinds and number of oribatid mites, whereas there are few such
studies covering the cool-temperate region. Yamashita and Ishii (1976) investigated arboreal arthropods
in the Daitsetsu mountain range in central Hokkaido, and recorded a high percentage of Acari in their
collected total. As they collected samples using canopy fog
canopy.
(3) Di百erenceof total organic matter: The total amount of organic matter is the most important fac-
tor influencing the abundance and organization of the Collembola and Oribatida communities (Hasegawa,
2001; Fagan et al., 2006). Lower diversity and abundance in bark and canopy than found in the forest
floor may be due to a smaller amount of litter and other organic matter. The trunk surface of oak from
which some oribatids were collected contains a small amount of suspended soil and other organic matter
in fissures and crevices, while the surface of leaves have few or no organic matter available as the food
for most oribatids. The fact that no oribatids were collected from leaves in our present study may be as
a result of lower total organic matter. However, further study, and a detailed comparable investigation on
organic matter in each site in particular, will be necessary to determine this.
We found the vertical distribution of canopy mites to be di旺erentamong each species. For the phy-
33
高久 元・佐々木富也
toseiid mites collected from canopy leaves, Euseius fi・nlandicuswas distributed mainly in lower positions
(5m and 10m), while Tyρhlodromus vulgaris was collected in a higher position (15m). Toyoshima and
Amano (2006), investigating the vertical distribution of predatory phytoseiid mites (such as E.β:nlandi-
cus, T. vulgaris, Amblyseius orientalis) on leaves at heights of 2m and 5m on three magnolia trees,
showed that the mite number did not di妊'erbetween higher and lower positions, but was significantly dif-
ferent between trees. T. vulgaris was collected from only one tree, and occurred in low numbers. As
sampling from a much higher position of the tree canopy was not carried out in their study, it's difficult
to compare their results with those of our current study. Winchester (1997) noted that tree height did
not affect guild proportionality, and predators, comprising mainly web-constructing arachnids, seemed
able to utilize the entire vertical. profile of the canopy. To confirm whether the vertical distribution of
predatory phytoseiid mitesshown in the present study is an exceptional case, mite distribution must be
investigated on several oak trees at a range of di旺erentheights.
E. finlandicus has been recorded in Japan in both Quercus mongolica var. grosseserrata and magnolia
(Ehara, 1958; Toyoshima and Amano, 2006), and T. vulgaris has been found in a ,variety of different
plants, such as citrus, plum, apple, willow, fir, and sasa bamboo (Ehara, 1959, 1962, 1964). These phy-
toseiid species prey on spider mites or eriophyid mites (Ehara, 1966). .According to Ehara and Ohashi
(2005),7 species of spider mites have so far been recorded as being found in the oak, Quercus spp., in
J apan. Phytoseiid mites collected from leaves in the oak canopy probably prey on spider mites or
eriophyids that inhabit leaves, and the distribution of phytoseiids in the canopy may depend on the dis-
tribution of prey. We have still not identified the mites of Prostigmata collected on leaves and bark of the
oak, however, and any correlation between the distribution of phytoseiids and spider mites so far re-
mains unknown.
The oribatid mites collected from the bark surface also exhibited a species-specific distributional pat-
tern: Ceratoρρia sp. was found to be distribl:lted on the bark in lower parts of tree, whereas Hexachaeto-
niella japonicus and Eρoribatula sp. near tuberosa were found on the bark in higher parts of tree. Wun-
derle (1992) has shown that oribatid species are usually dominant at particular levels of trunk and
canopy. The distribution and species richness of oribatid mites on bark is related to the type of epiphytic
cover (Behan-Pelletier and Walter, 2000). Nicolai (1986) found that highly diverse fauna lived on飴sured
bark, although Beaulieu et al. (2006) concluded that bark roughness had no significant e旺ecton abund-
ance or species richness of mesostigmatic mites. Any specificity to certain heights found in the present
study may be caused by distribution of organic matter (such as litter, fungi, moss, and lichen), bark
structure Cincluding such as smooth, fissure, and crevice), or abiotic factors Cincluding temperature,
moisture, desiccation, and insolation) on bark surface, although further study remains necessary.
ACKNOWLEDGEMENTS
We would like to express our sincere thanks to Drs. M. J. Toda, T. Hiura, and M. Murakami
(Hokkaido University) for giving us the opportunity to study this subject and for use of facilities in the
Tomakomai Experimental Forest of Hokkaido University. This study was partly supported by Grant-in-
Aid for Scientific Research (Nos. 15207008, 18570078) from the Japan Society for the Promotion of Scien-
34
北海道大学苫小牧研究林tこ見られる樹上性および土壊性ダニ類 (ダニ目 トゲダニ亜日,ササラダニ亜日)
ce (JSPS) and a President Grant in Aid for Research (Special Research Grant for Facilities) from Hok-
kaido University of Education.
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(高久 元札幌校准教授)
(佐々木富也札幌市立和光小学校教諭)
36
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